Polish Botanical Journal 59(1): 85–96, 2014 DOI: 10.2478/pbj-2014-0017

A CONTRIBUTION TO THE FAMILY (, ) OF BOLIVIA. 2

Ulf Schiefelbein, Adam Flakus, Harrie J. M. Sipman, Magdalena Oset & Martin Kukwa1

Abstract. Microlichens of the family Graphidaceae are important components of the lowland and montane tropical forests in Bolivia. In this paper we present new records for 51 taxa of the family in Bolivia. Leiorreuma lyellii (Sm.) Staiger is reported as new for the Southern Hemisphere, while caesioplumbeus (Nyl.) Vain., daintreensis (A. W. Archer) A. W. Archer, G. duplicatoinspersa Lücking, G. emersa Müll. Arg., G. hossei Vain., G. immersella Müll. Arg. and G. subchrysocarpa Lücking are new for South America. Thirty taxa are reported for the first time from Bolivia. Notes on distribution are provided for most species. Key words: biodiversity, biogeography, lichenized fungi, Neotropics, South America Ulf Schiefelbein, Blücherstr. 71, D-18055 Rostock, Germany Adam Flakus, Laboratory of Lichenology, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland Harrie J. M. Sipman, Botanischer Garten & Botanisches Museum Berlin Dahlem, Königin-Luise-Strasse 6 – 8, D-14195 Berlin, Germany Martin Kukwa & Magdalena Oset, Department of Plant and Nature Conservation, University of Gdańsk, Wita Stwosza 59, 80-308 Gdańsk, Poland; e-mail: [email protected]

Introduction

Bolivia has the highest ecosystem diversity in Material and methods South America and the forest communities form a mosaic of vegetation which offers a variety Specimens are deposited at B, GOET, KRAM, LPB, of potential habitats for numerous microlichens UGDA (acronyms after Thiers 2012) and the private (Navarro & Maldonado 2002; Josse et al. 2003; herbaria of A. Flakus and U. Schiefelbein. Secondary Ibisch & Mérida 2004; Navarro & Ferreira 2007). lichen metabolites were identified by thin layer chro- matography (TLC) according to Orange et al. (2001). Thus a large number of such species should be Data on general distribution are presented only for expected to occur in Bolivia; however, current species reported for the first time from Bolivia. New knowledge of this group in Bolivia is very in- national records are asterisked (*), those new for South complete, and the family Graphidaceae is no America have two asterisks (**), and those new for the exception in this regard. On the basis of mate- Southern Hemisphere have three (***). rial we collected recently during lichenological The following abbreviations are used in the “Speci- exploration in Bolivia, here we document some mens examined” sections: AF – A. Flakus; MK – M. of this hidden diversity. Kukwa; OP – O. Plata; PR – P. Rodriguez Flakus; RIM This paper is the second contribution dealing – R. I. Meneses; US – U. Schiefelbein; RN – Reserva Nacional; AMBORÓ – Parque Nacional y Área Natural with the family Graphidaceae in Bolivia (Kukwa de Manejo Integrado Amboró; APOLOBAMBA – Área et al. 2013). Natural de Manejo Integrado Nacional Apolobamba; CARRASCO – Parque Nacional Carrasco; COTAPATA – Parque Nacional y Área Natural de Manejo Integrado 1 Corresponding author Cotapata; KAA-IYA – Parque Nacional y Área Natural 86 POLISH BOTANICAL JOURNAL 59(1). 2014 de Manejo Integrado Kaa-Iya del Gran Chaco; SAJAMA epiglaucum (Müll. Arg.) Kalb, Staiger – Parque Nacional Sajama; TARIQUÍA – Reserva Na- & Elix cional de Flora y Fauna Tariquía (abbreviations of pro- tected areas according to SERNAP Bolivia; http://www. Previously known from three records in Bolivia sernap.gob.bo). (Kalb et al. 2004; Flakus 2008). Specimens examined. BOLIVIA. Dept. Beni. Results Prov. Marbán, near road from Camiaco to San Lor- enzo, 15°14′49″S, 64°49′11″W, 160 m, lowland *Diorygma antillarum (Vain.) Nelsen, Lücking Amazon forest surrounded by los Llanos de Moxos & Rivas Plata savanna, corticolous, 30 Aug. 2012, MK 11951 (LPB, UGDA); Dept. Santa Cruz. Prov. Guarayos, RN de A Neotropical species previously reported Vida Silvestre Ríos Blanco y Negro, Virgen de Pilar from Brazil, Costa Rica, El Salvador, the Lesser near Chonta, 15°38′54″S, 62°57′37″W, 229 m, lowland Antilles, Panama, Peru, Venezuela and the USA Amazon forest, corticolous, 22 July 2009, AF & PR (Florida) (Aptroot et al. 2009; Nelsen et al. 2012). 13143 (KRAM, LPB).

Specimens examined. BOLIVIA. Dept. Tarija. Prov. Diorygma intermedium Kalb, Staiger & Elix Aniceto Arce, Filo de Sidras, 22°14′50″S, 64°33′28″W, 1064 m, Tucumano-Boliviano submontane forest, on Kukwa et al. (2013) reported this species from bark, 22 Nov. 2010. AF 18421 & 18672 (KRAM, LPB); four localities in Bolivia. Serranía de Propiedad Arnold, 22°13′19″S, 64°33′41″W, 1309 m, Tucumano-Boliviano montane forest, on Specimens examined. BOLIVIA. Dept. Santa bark, 24 Nov. 2010, AF 18708 (KRAM, LPB); Prov. Cruz. Prov. Guarayos, RN de Vida Silvestre Ríos ° ′ ″ Burnet O’Connor, Lomas de la Soledad, road between Blanco y Negro, Plan de Manejo AISU, 15 01 49 S, ° ′ ″ Entre Ríos and Chiquiacá, 21°39′38″S, 64°07′31″W, 62 46 36 W, 236 m, lowland Amazon forest, corti- 1670 m, Tucumano-Boliviano altimontane forest, cor- colous, 25 July 2009, AF 14200 & PR (KRAM, LPB). ticolous, 10 Aug. 2012, MK 11264 (LPB, UGDA); TARIQUÍA, near Salinas, 21°49′15″S, 64°12′44″W, Diorygma poitaei (Fée) Kalb, Staiger & Elix 1400 m, Tucumano-Boliviano montane forest, corti- This Neotropical lichen was recently reported colous, 10 Aug. 2012, MK 11246 (LPB, UGDA). from two localities in Bolivia (Kukwa et al. 2013). Diorygma confluens (Fée) Kalb, Staiger & Elix Specimens examined. BOLIVIA. Dept. Beni. Prov. Cercado, Casa del Tigre near Chuchini, 14°43′17″S, This species was only recently reported from 64°56′53″W, 160 m, lowland Amazon forest, corti- Bolivia from six localities (Kukwa et al. 2013). colous, 31 Aug. 2012, MK 11997 (LPB, UGDA). Diorygma confluens is chemically variable (Kalb et al. 2004). The material reported so far *Diorygma reniforme (Fée) Kalb, Staiger & Elix from Bolivia contained lichexanthone, stictic and This species has been reported from the Neo- constictic acids (major), and sometimes traces of tropics in Brazil, Colombia, Cuba, British Guiana, hypoconstictic, hypostictic, norstictic and cryp- Peru and Venezuela, and elsewhere from Cam- tostictic acids (Kukwa et al. 2013). In one of the eroon and Tanzania (Kalb et al. 2004). specimens cited below (MK 7111), lichexanthone, hypoconstictic (major), constictic (submajor) and Specimen examined. BOLIVIA. Dept. Santa Cruz. stictic (trace) acids were detected. Prov. Guarayos, RN de Vida Silvestre Ríos Blanco y Negro, Plan de Manejo AISU, 15°01′58″S, 62°46′36″W, Specimens examined. BOLIVIA. Dept. Beni. Prov. 242 m, lowland Amazon forest, corticolous, 24 July Ballivian, Palmar, 14°58′56″S, 67°05′16″W, 250 m, by 2009, AF 13899 & PR (LPB). road, corticolous, 2 Aug. 2008, MK 7111 (LPB, UGDA); Dept. La Paz. Prov. Iturralde, between Tumapasa and *Diorygma sipmanii Kalb, Staiger & Elix Mamuque, 300 m, garden, corticolous, 5 Mar. 1987, S. Stab LB-33 (B). This Neotropical species was known previ- U. SCHIEFELBEIN ET AL.: A CONTRIBUTION TO GRAPHIDACEAE OF BOLIVIA. 2 87 ously from Brazil, Costa Rica, Guatemala and El ceous schist, 31 May 2011, AF & OP 22123 (KRAM, Salvador (Kalb et al. 2004, Cáceres 2007). LPB); near Cumbre pass, 16°19′18″S, 68°04′42″W, 4550 m, high Andean Puna, on siliceous rock, 17 June Specimen examined. BOLIVIA. Dept. Santa Cruz. 2006, AF 8530 (herb. Flakus). Prov. Guarayos, RN de Vida Silvestre Ríos Blanco y Negro, Virgen de Pilar near Chonta, 15°38′54″S, Diploschistes cinereocaesius (Sw. ex Ach.) Vain. 62°57′37″W, 229 m, lowland Amazon forest, corti- colous, 22 July 2009, AF 13273 & PR (LPB). This species was reported from Bolivia by Feuerer et al. (1998) and Flakus et al. (2012). It Diploschistes actinostomus (Ach.) Zahlbr. appears to be a common lichen at high elevations in the Andes. This species was reported in Bolivia from only one locality (Feuerer & Sipman 2005) Specimens examined. BOLIVIA. Dept. Cocha- bamba. Prov. Quillacollo, area of Incarraya-Sipesipe, Specimens examined. BOLIVIA. Dept. La Paz. 17°29′25″S, 66°22′09″W, 3146 m, semi-desert open Prov. Camacho, Pacoamba cerca Wila Kala, 15°24′40″S, area, on soil, 17 Dec. 2004, AF 4977 (LPB); Dept. La 69°04′24″W, 4283 m, Puna Húmeda, on sandstone, Paz. Prov. Camacho, cerca de la comunidad de Ulla 6 July 2010, AF 17740 & PR (KRAM); Prov. Manco Ulla, en el cerro Wilamuku, 15°02′04″S, 69°11′53″W, Kapac, near Copacabana Mt. Horca del Inca, 16°10′15″S, 4739 m, vegetación altoandina, terrestre, 4 Dec. 69°05′05″W, 3974 m, high Andean Puna, on volcanic 2007, PR 272 (B, LPB); near Villa Cala, 15°25′53″S, rock, 18 June 2006, AF 8637 (LPB). 69°05′07″W, 4250 m, high Andean vegetation, on soil, 19 May 2011, AF 21236 & OP (KRAM, LPB); Prov. **Diploschistes caesioplumbeus (Nyl.) Vain. Franz Tamayo, APOLOBAMBA, Socondori Chico near Ulla Ulla, 15°00′38″S, 69°13′48″W, 4479 m, high An- This species was known previously from Af- dean open vegetation, on soil, 4 July 2010, AF 17473 rica (e.g., Algeria, Canary Islands, Madeira, Mo- & PR (KRAM, LPB); Prov. Murillo, near Cumbre pass, rocco, St. Helena and Ascension Island), Asia (e.g., 16°19′18″S, 68°04′42″W, 4550 m, high Andean Puna, on India, Israel, Turkey), Europe (the Azores, France, soil and terricolous bryophytes, 17 June 2006, AF 8533 Greece, Great Britain, Ireland, Italy, Spain) and (KRAM, LPB, herb. Flakus); Valle del Zongo, Laguna western North America (Breuss 1988; Lumbsch Viscachani, 16°11′54″S, 68°07′33″W, 3862 m, Páramo 1989, 2002; Follmann 1990; Pant & Upreti 1993; Yungueño, on soil, 30 May 2011, AF 22085, 22103 & OP (KRAM, LPB); Prov. Sud Yungas, Mururata, Hafellner 1995; Llimona & Hladun 2001; Yazici bajo la Mina Bolsa Negra, 16°35′S, 67°45′W, 4000 m, & Aslan 2003; Aptroot 2008; Seaward 2010; Roux saxicolous, 1995, J. Gonzáles 106 (B, LPB); ibidem, 2012; Esslinger 2014). pasando por el pueblo de Tres Rios, 16°35′S, 67°45′W, 3851 m, saxicolous, 6 Apr. 1995, J. Gonzáles 380 (B, Specimens examined. BOLIVIA. Dept. La Paz. LPB); Dept. Santa Cruz. Prov. Manuel María Cabal- Prov. Bautista Saavedra, APOLOBAMBA, near Villa lero, Siberia, 17°49′38″S, 64°45′14″W, 3000 m, open Amarca, 15°16′47″S, 69°01′47″W, 4643 m, Puna area near Yungas cloud forest, on soil, 11 Dec. 2004, AF Húmeda, on sandstone, 3 July 2010, AF 17349, 17355 4474 & 4478 (LPB, herb. Flakus); Dept. Tarija. Prov. & PR (KRAM, LPB); Prov. Camacho, Pacoamba Méndez, Cuesta de Sama, near Las Antenas, 21°29′59″S, cerca Wila Kala, 15°24′40″S, 69°04′24″W, 4283 m, 64°54′46″W, 3800 m, high Andean vegetation, 6 Aug. Puna Húmeda, on sandstone, 6 July 2010, AF 17708, 2012, MK 10958 (LPB, UGDA). 17727, 17757, 17769, 17788 & PR (KRAM, LPB, herb. Flakus); Prov. Franz Tamayo, APOLOBAMBA, near * Puyo Puyo, 14°56′55″S, 69°07′58″W, 4888 m, high Diploschistes diacapsis (Ach.) Lumbsch Andean open vegetation, on siliceous schist, 5 July 2010, This is a widely distributed species known from AF 17564 & PR (KRAM, LPB); Prov. Murillo, below Potosí, on road between La Paz and Valle del Zongo, Africa (e.g., Algeria, Canary Islands, Morocco, Tu- 16°17′43″S, 68°07′42″W, 4716 m, high Andean vegeta- nisia), Asia (e.g., China, Cyprus, India, Iran, Mon- tion, on granite rock, 28 May 2011, AF 21840, 21860 golia, Saudi Arabia, Syria), Australia, Europe (e.g., & OP (KRAM, LPB); near Cumbre pass, 16°19′06″S, Albania, Azores, Bosnia and Herzegovina, France, 68°02′09″W, 4405 m, high Andean vegetation, on sili- Germany, Greece, Spain), North America (Mexico, 88 POLISH BOTANICAL JOURNAL 59(1). 2014

USA) and South America (Chile, Colombia, Peru) and elsewhere in Oceania (Hawaii), continental (Schubert & Klement 1971; Lumbsch 1989; Pant North America, the Caribbean (Puerto Rico) and & Upreti 1993; Hafellner 1995, 2007; Galloway the Republic of South Africa (Lumbsch 1989; & Quilhot 1998; Litterski & Mayrhofer 1998; Lli- Guderley & Lumbsch 1996; Marcano et al. 1996; mona & Hladun 2001; Scholz 2001; John et al. Elix & McCarthy 1998; Esslinger 2014). 2004; Seaward et al. 2008; Sipman et al. 2008; Specimens examined. BOLIVIA. Dept. Cocha- Bilovitz & Mayrhofer 2011; Roux 2012; McCarthy bamba. Prov. Carrasco, CARRASCO, La Cumbre, El 2013; Esslinger 2014). Camino de las Nubes, 17°17′46″S, 65°43′56″W, 4100 m, Specimens examined. BOLIVIA. Dept. La Paz. Páramo Yungueño, on soil, 19 Aug. 2012, MK 11806 Prov. Omasuyos, El Dragon hill near Chahualla, (LPB, UGDA); Prov. Quillacollo, area of Incarraya- ′ ″ ′ ″ 15°51′17″S, 69°00′40″W, 3850 m, Puna Húmeda, on Sipesipe, 17°29 25 S, 66°22 09 W, 3146 m, semi-desert siliceous rock, 6 July 2010, AF 17836 & PR (KRAM, open area, on soil, 17 Dec. 2004, AF 4964 (KRAM, LPB, herb. Flakus). LPB, herb. Flakus); Dept. La Paz. Prov. Larecaja, la Cumbre de Sorata, 15°53′06″S, 68°38′55″W, 4210 m,

* high Andean open vegetation, on soil, 13 May 2011, Diploschistes diploschistoides (Vain.) G. Salisb. AF 20256 & OP (KRAM, LPB). This species was reported previously from * South Africa, Zimbabwe, Australia, and in South (Scop.) R. Sant. subsp. America from Brazil and Colombia (Guderley bartletii Lumbsch & Lumbsch 1996; McCarthy 2013). This taxon is less common than D. muscorum Specimens examined. BOLIVIA. Dept. Tarija. subsp. muscorum and has a remarkable Gondwa- Prov. Aniceto Arce, Papachacra, near Papachacra valley, naland distribution. It was known previously from 21°41′14″S, 64°30′19″W, 2050 m, open vegetation with Australia, India, Lesotho, the Republic of South Af- shrubs, on sandstone, 7 Aug. 2012, AF 23911 (LPB, rica, and in South America from Argentina, Brazil, herb. Flakus). Chile, Colombia and Ecuador (Galapagos Islands) (Pant & Upreti 1993; Guderley & Lumbsch 1996; *Diploschistes euganeus (A. Massal.) Zahlbr. Elix & McCarthy 1998; Galloway & Quilhot 1998; Aptroot 2002; Calvelo & Liberatore 2002; Sipman Reported in South America only from Brazil et al. 2008; McCarthy 2013). and Chile, and elsewhere from Africa (Algeria, Cape Verde, Republic of South Africa, St. He- Specimens examined. BOLIVIA. Dept. La Paz. lena), Asia (Taiwan), Australia (mainland Aus- Prov. Omasuyos, El Dragon hill near Chahualla, tralia, New Zealand) and Europe (France, Greece, 15°51′17″S, 69°00′40″W, 3850 m, Puna Húmeda, on Italy, Poland, Spain) (Lumbsch 1989; Mies soil, 6 July 2010, AF 17837.1 & PR (KRAM, LPB); Dept. La Paz. Prov. Nor Yungas, near Pongo, near the & Lumbsch 1990; Guderley & Lumbsch 1996; Coroico-La Paz road, 16°19′28″S, 67°57′21″W, 3822 m, Galloway & Quilhot 1998; Llimona & Hladun Páramo Yungueño, on soil, 25 May 2011, AF 21736 2001; Fałtynowicz 2003; Aptroot 2008; Roux & OP (LPB). 2012; McCarthy 2013). * Specimens examined. BOLIVIA. Dept. Cocha- Diploschistes muscorum (Scop.) R. Sant. subsp. bamba. Prov. Quillacollo, area of Incarraya-Sipesipe, muscorum ′ ″ ′ ″ 17°28 39 S, 66°21 43 W, 2846 m, semi-desert open This widely distributed taxon is known from area, on sandstone, 17 Dec. 2004, AF 4922 (LPB, herb. Flakus). Africa (Cape Verde), Asia (India, Iran), Europe (e.g., Bosnia and Herzegovina, Finland, France, Norway, Poland, Sweden), North America and *Diploschistes hypoleucus Zahlbr. South America (Argentina, Chile, Colombia, Ec- This rarely reported species is known from uador) (Mies & Lumbsch 1990; Pant & Upreti Brazil, Peru and Venezuela in South America, 1993; Elix & McCarthy 1998; Calvelo & Lib- U. SCHIEFELBEIN ET AL.: A CONTRIBUTION TO GRAPHIDACEAE OF BOLIVIA. 2 89 eratore 2002; Fałtynowicz 2003; Santesson et al. *Graphis caesiella Vain. 2004; Nöske et al. 2007; Seaward et al. 2008; This is a pantropical species reported in the Neo- Sipman et al. 2008; Bilovitz & Mayrhofer 2011; tropics from Brazil, Peru and the USA (Louisiana, Roux 2012; Esslinger 2014). Florida) (Staiger 2002; Cáceres 2007; Lücking et al. Specimens examined. BOLIVIA. Dept. La Paz. 2009; Seavey & Seavey 2011; Rivas Plata & Lücking Prov. Bautista Saavedra, km 162 on road from Apolo 2013), and also from the Philippines (Linsangan- to Charazani, la Cruz Charazani-Pelechuco, 15°15′00″S, Tabaquero et al. 2013) and Australia (Archer 2009). 69°02′51″W, 4545 m, high Andean vegetation, on soil, 19 May 2011, AF 21160 & OP (LPB); Dept. Oruro. Specimen examined. BOLIVIA. Dept. Santa SAJAMA, Ladera de la Quebrada Kohniri. 18°07′07″S, Cruz. Prov. Ichilo, AMBORÓ, Macuñucu, 17°43′38″S, 68°56′06″W, 4309 m, Polylepis tarapacana forest, on 63°35′37″W, 460 m, Amazon forest, on twigs, 6 June rock, 21 Nov. 2007, RIM et al. 4380, 4385 (B, LPB); 2011, MK 9696a (LPB). ibidem, 18°07′08″S, 68°56′05″W, 4406 m, on rock, 21 Nov. 2007, RIM et al. 4391 (B, LPB); Prov. Sajama, SA- *Graphis cincta (Pers.) Aptroot JAMA, Jecha K’ala 18°09′52″S, 68°49′08″W, 4184 m, Puna Sureña, on soil, 20 June 2010, AF 16679.1, 16684 This species was previously reported in the & PR (KRAM, LPB). Neotropics from Costa Rica, the Dominican Re- public, Paraguay, the USA (Florida), and elsewhere afzeli (Ach.) A. Massal. from the Philippines and Taiwan (Lücking et al. 2008, 2009; Seavey & Seavey 2011) and from This easily recognizable species is perhaps Australia and Japan (Archer 2009). the commonest member of the Graphidaceae in Bolivia. It was previously reported from there by Specimens examined. BOLIVIA. Dept. La Paz. Kalb & Staiger (2000), Staiger (2002), Flakus Prov. Sud Yungas, Alto Beni, San Antonio. 15°38′13″S, ° ′ & Wilk (2006), Flakus (2008) and Flakus et al. 67 12 09″W, 405 m, on Theobroma cacao, 29 June 1999, N. Derakshani 12 (B, GOET). (2012). Specimens examined. BOLIVIA. Dept. La Paz. **Graphis daintreensis (A. W. Archer) A. W. Archer Prov. Iturralde, near San Pedro, 13°43′59″S, 68°00′38″W, 250 m, savannah, corticolous, 28 July 2008, MK 6897 Previously known only from Australia and the (LPB, UGDA); Tumupasa, 14°08′51″S, 67°53′34″W, Solomon Islands (Archer 2009). 350 m, garden, on mango tree, 31 Aug. 2008, MK 6996 Specimens examined. BOLIVIA. Dept. Beni. (LPB, UGDA). Prov. Vaca Díez, Cachuela Esperanza, 10°32′09″S, 65°34′55″W, 128 m, lowland Amazon forest, corti- * triticea (Nyl.) Staiger colous, 4 June 2006, AF 7525 (KRAM, LPB).

This species was previously reported from Aus- * tralia, Madeira, New Zealand, Reunion, the Azores, Graphis dracaenae Vain. and in the Neotropics from Colombia, Costa Rica, Previously known from Africa, Australia and Dominica, Jamaica and Mexico (Wirth & Hale Brazil (Archer 2009; Lücking et al. 2009). 1978; Hafellner 1995; Staiger 2002; Sipman 2006; Specimen examined. BOLIVIA. Dept. La Paz. Sipman et al. 2008). Prov. Nor Yungas, COTAPATA, Santa Catalina, above Specimens examined. BOLIVIA. Dept. La Tunkini, 16°11′12″S, 67°52′07″W, Yungas montane Paz. Prov. Nor Yungas, near Nogalani, 16°12′57″S, forest, corticolous, 22 May 2011, US (LPB). 67°49′15″W, 2168 m, Yungas secondary cloud forest, corticolous, 23 May 2011, US (LPB, herb. Schiefel- Graphis duplicata Ach. bein 3069, 3070); Dept. Santa Cruz. Prov. Manuel María Caballero, El Camino de Orquideas, 17°49′20″S, Reported only recently from three localities in 64°42′31″W, 2340 m, Yungas cloud forest, corticolous, Bolivia by Flakus et al. (2013) and Kukwa et al. 17 Aug. 2012, MK 11564 (LPB, UGDA). (2103). 90 POLISH BOTANICAL JOURNAL 59(1). 2014

Specimen examined. BOLIVIA. Dept. Santa Cruz. *Graphis handelii Zahlbr. Prov. Manuel María Caballero, Siberia region near La Palma, 17°49′12″S, 64°40′28″W, 2582 m, Yungas cloud Previously reported from Brazil, China, Costa forest, corticolous, 13 Dec. 2004, AF 4642 (KRAM, Rica, India and Paraguay (Lücking et al. 2008, LPB). 2009). Specimens examined. BOLIVIA. Dept. La Paz. ** Graphis duplicatoinspersa Lücking Prov. Nor Yungas, between Coroico and La Paz, ° ′ ″ ° ′ ″ The species has been reported from Costa Rica 16 13 33 S, 67 49 20 W, 2600 m, Yungas secondary cloud forest, corticolous, 23 May 2011, US (LBP); and Australia (Lücking et al. 2008, 2009). Dept. Santa Cruz. Prov. Guarayos, RN de Vida Sil- Specimen examined. BOLIVIA. Dept. La Paz. vestre Ríos Blanco y Negro, Plan de Manejo AISU, Prov. Nor Yungas, COTAPATA, near Urpuma colony, 15°01′49″S, 62°46′36″W, 236 m, lowland Amazon 16°13′20″S, 67°52′34″W, 1989 m, Yungas montane forest, corticolous, 25 July 2009, AF 14254 & PR forest, corticolous, 30 June 2010, AF 17118 & PR (LPB). (KRAM, LPB, herb. Flakus). **Graphis hossei Vain. ** Graphis emersa Müll. Arg. Previously this species was known only from Previously reported from Australia, Costa Rica Asia (China, Indonesia, Philippines and Thailand) and Japan (Lücking et al. 2008, 2009). (Lücking et al. 2009). Specimen examined. BOLIVIA. Dept. La Paz. Prov. Specimen examined. BOLIVIA. Dept. La Paz. Nor Yungas, between Coroico and La Paz, 16°13′33″S, Prov. Nor Yungas, COTAPATA, Chairo, 16°12′07″S, 67°49′20″W, 2600 m, Yungas secondary cloud forest, 67°50′05″W, 1300 m, Yungas lower montane forest, corticolous, 23 May 2011, US (LBP). corticolous, 21 May 2011, US (LPB).

Graphis aff. evirescens (Redinger) Lücking Graphis illinata Eschw. To our knowledge this species was previously Reported from Bolivia by Flakus & Wilk reported only from the type locality in Brazil (2006). (Lücking et al. 2009). Specimens examined. BOLIVIA. Dept. La Paz. With Lücking et al. (2009), our material keys Prov. Nor Yungas, near Nogalani, between Coroico out as G. evirescens, characterized by a green and La Paz, 16°12′57″S, 67°49′15″W, 2168 m, Yungas thallus, stellately branched lirellae with an api- secondary cloud forest, corticolous, 23 May 2011, US cally thin, complete thalline margin, 8-spored (LPB, herb. Schiefelbein 3453); Prov. Nor Yungas, CO- asci and small muriform ascospores, 12–18 × TAPATA, Santa Catalina, above Tunkini, 16°11′12″S, 67°52′07″W, Yungas montane forest, corticolous, 7–14 µm (Redinger 1935; Lücking et al. 2009). 22 May 2011, US (LPB, herb. Schiefelbein 3057). In our specimen the lirellae are unbranched, the asci 2-spored and the ascospores larger (30–44 × **Graphis immersella Müll. Arg. 13–17 µm). It is not clear to us if this should be considered an instance of infraspecific variation, Known previously from Australia, China, the such as known in other Graphis species (Lücking Philippines, New Caledonia, the Solomon Islands et al. 2009), so the present report should be con- and Vanuatu (Archer 2009; Lücking et al. 2009; sidered tentative. Linsangan-Tabaquero et al. 2013). Specimen examined. BOLIVIA. Dept. La Paz. Specimens examined. BOLIVIA. Dept. Santa Prov. Nor Yungas, COTAPATA, between Tunkini and Cruz. Prov. Guarayos, RN de Vida Silvestre Ríos Blanco Chairo villages, above Tunkini, 16°11′S, 67°52′W, y Negro, Virgen de Pilar near Chonta, 15°38′54″S, 2400 m, Yungas montane forest, corticolous, 22 May 62°57′37″W, 229 m, lowland Amazon forest, corti- 2011, MK 9101 (LPB, UGDA). colous, 22 July 2009, AF 13217 & PR (KRAM, LPB). U. SCHIEFELBEIN ET AL.: A CONTRIBUTION TO GRAPHIDACEAE OF BOLIVIA. 2 91

*Graphis librata Vain. 2340 m, Yungas cloud forest, corticolous, 17 Aug. 2012, MK 11565 (LPB, UGDA). In the Neotropics this species was found in Brazil, Costa Rica, Panama and Venezuela, and *Graphis paraserpens Lizano & Lücking elsewhere in Australia, New Zealand, the Philip- pines, South Africa and the USA (Archer 2001; So far reported from Brazil and Costa Rica Staiger 2002; Lücking et al. 2008, 2009; Neuwirth (Cáceres 2007; Lücking et al. 2008, 2009). 2008; Seavey & Seavey 2011; Boom & Sipman Specimens examined. BOLIVIA. Dept. Santa 2013; Linsangan-Tabaquero et al. 2013). Cruz. Prov. Guarayos, RN de Vida Silvestre Ríos Blanco y Negro, Plan de Manejo AISU, 15°01′49″S, Specimens examined. BOLIVIA. Dept. Santa 62°46′36″W, 236 m, lowland Amazon forest, corti- Cruz. Prov. Cordillera, KAA-IYA, near Peto Blanco, colous, 25 July 2009, AF 14250 & PR (KRAM, LPB). 18°56′26″S, 60°22′39″W, 340 m, Chiquitano forest, on twig, 5 Dec. 2011, MK 10644 (LPB, UGDA). *Graphis phaeospora Vain. Graphis lumbricina Vain. So far known only from Brazil and Ecuador (Staiger 2002; Nöske et al. 2007). Previously reported from four localities in Bolivia (Flakus et al. 2103; Kukwa et al. 2013). Specimens examined. BOLIVIA. Dept. Cocha- bamba. Prov. Carrasco, CARRASCO, near Sehuencas, Specimens examined. BOLIVIA. Dept. Santa 17°30′12″S, 65°16′30″W, 2220 m, montane cloud forest, Cruz. Prov. Manuel María Caballero, near Siberia, along road, corticolous, 21 July 2008, MK 6395 (LPB, 17°49′38″S, 64°44′45″W, 3950 m, open Yungas cloud UGDA); Dept. Santa Cruz. Prov. Manuel María Ca- forest, corticolous, 16 Aug. 2012, MK 11477 (LPB, ballero, Siberia region near La Palma, 17°49′12″S, UGDA). 64°40′28″W, 2582 m, Yungas cloud forest, on sand- stone, 13 Dec. 2004, AF 4641 (LPB). *Graphis paralleloides Cáceres & Lücking *Graphis pitmanii Rivas Plata & Lücking Reported from Brazil and India (Cáceres 2007; Lücking et al. 2009). This species was only recently described from According to Cáceres (2007) and Lücking et al. Peru (Rivas Plata & Lücking 2013). (2009) this species produces transversely septate Specimens examined. BOLIVIA. Dept. La Paz. Prov. ascospores measuring 30–40 × 7–8 µm. In one Bautista Saavedra, Calzada, 15°09′06″S, 68°30′24″W, collections cited below (MK 11565) the ascospores 834 m, Preandean Amazon secondary forest, corticolous, were 38–50 × 13–14 µm, and in the other (MK 17 May 2011, US (LPB, herb. Schiefelbein 2012); Prov. 9792a) 25–40 × 7–12 µm, but otherwise both Larecaja, near Achiquiri, 15°13′31″S, 68°13′49″W, collection had the same morphology of ascomata 1071 m, Yungas secondary forest, corticolous, 16 May (striatula-morph) as presented by Cáceres (2007) 2011, US (LPB, herb. Schiefelbein 2988). and Lücking et al. (2009). Considerable variation * of ascospore dimensions is known in other Graphis Graphis puiggarii (Müll. Arg.) Lücking species as well (e.g., in Graphis conferta Zenker Known before from Brazil, Costa Rica and or G. ovata (Fée) A. Massal.; Lücking et al. 2009), Paraguay (Lücking et al. 2008, 2009). so here we widen the circumscription of G. paral- leloides instead of distinguishing a new species. Specimens examined. BOLIVIA. Dept. La Paz. Prov. Franz Tamayo, near Correo, 14°59′06″S, Specimens examined. BOLIVIA. Dept. Santa 68°26′09″W, 1632 m, Yungas montane forest, open area, Cruz. Prov. Florida, AMBORÓ, above la Yunga vil- corticolous, 17 May 2011, US (LPB, herb. Schiefel- lage, senda Los Helechos, 18°03′30″S, 63°54′36″W, bein 3002); Prov. Larecaja, near Incachaca, 15°17′48″S, 2330 m, Yungas cloud forest, on dead twig, 7 June 68°26′32″W, 1100 m, Yungas lower montane forest, 2011, MK 9792a (LPB); Prov. Manuel María Cabal- corticolous, 15 May 2011, US (LPB, herb. Schiefelbein lero, El Camino de Orquideas, 17°49′20″S, 64°42′31″W, 2980); Prov. Nor Yungas, COTAPATA, Santa Catalina, 92 POLISH BOTANICAL JOURNAL 59(1). 2014 above Tunkini, 16°11′12″S, 67°52′07″W, Yungas mon- Prov. Nor Yungas, COTAPATA, between Tunkini and tane forest, 22 May 2011, corticolous, US (LPB, herb. Chairo, 16°11′00″S, 67°52′00″W, corticolous, 22 May Schiefelbein 3029); Chairo, 16°12′07″S, 67°50′05″W, 2011, US (LPB). Yungas lower montane forest, corticolous, 1300 m, 21 May 2011, US (LPB). *Graphis supracola A. W. Archer

*Graphis rhizocola (Fée) Lücking & Chaves A Pantropical species known from Australia, Cambodia, Japan and Thailand, and in the Ne- This Neotropical species was reported previ- otropics from Brazil, Colombia and the USA ously from Brazil, Costa Rica, Peru, St. Vincent (Florida) (Archer 2001, 2012; Lücking et al. 2009; and Trinidad (Lücking et al. 2008, 2009). Nakanishi et al. 2010; Seavey & Seavey 2011). Specimens examined. BOLIVIA. Dept. La Paz. Specimens examined. BOLIVIA. Dept. Beni. Prov. Prov. Nor Yungas, COTAPATA, Santa Catalina, above Marbán, near San Lorenzo, 15°12′59″S, 64°47′08″W, Tunkini, 16°11′12″S, 67°52′07″W, Yungas montane 160 m, lowland Amazon forest surrounded by los Llanos forest, 17 May 2011, US (LPB, herb. Schiefelbein 3056); de Moxos savanna, on twigs, 28 Aug. 2012, MK 11897 by Sillu Tincara pre-Columbian route, 16°17′22″S, (LPB, UGDA). 67°53′29″W, 3518 m, transition Páramo Yungeño– Yungas montane cloud forest, 25 May 2011, US (LPB, * laubertianum (Fée) Staiger herb. Schiefelbein 3456). This rare species is known from Dominica, *Graphis sitiana Vain. Peru and Australia (Staiger 2002; Archer 2009). A Neotropical species known previously from Specimens examined. BOLIVIA. Dept. Beni. Prov. ° ′ ″ Brazil, Costa Rica, Panama and Peru (Lücking Cercado, Casa del Tigre near Chuchini, 14 43 17 S, 64°56′53″W, 160 m, lowland Amazon forest, corti- et al. 2008, 2009; Boom & Sipman 2013; Rivas colous, 31 Aug. 2012, MK 11984 (LPB, UGDA, herb. Plata & Lücking 2013). Flakus). Specimens examined. BOLIVIA. Dept. La Paz. Prov. Sud Yungas, Alto Beni Sapecho. Cooperativa Sa- ***Leiorreuma lyellii (Sm.) Staiger pecho. 15°32′S, 67°20 ‘W, 500 m, on branches, 2 Aug. lyellii (Sm.) Zahlbr. 1999, A. Acebey & T. Krömer 997 (B, LPB). Previously reported from Great Britain, France, **Graphis subchrysocarpa Lücking Ireland, Spain and Macaronesia (Madeira), and in the Neotropics only from Costa Rica (Llimona This species is known from Central America & Hladun 2001, Staiger 2002; Breuss & Neuwirth (Costa Rica) and Africa (Lücking et al. 2008, 2007; Benfield et al. 2009; Seaward 2010; Roux 2009). 2012). Specimens examined. BOLIVIA. Dept. Santa Specimens examined. BOLIVIA. Dept. La Paz. Cruz. Prov. Guarayos, RN de Vida Silvestre Ríos Prov. Nor Yungas, near Chuspipata, 16°17′09″S, Blanco y Negro, Plan de Manejo AISU, 15°01′49″S, 67°51′00″W, 3228 m, Yungas cloud forest with Eri- 62°46′36″W, 236 m, lowland Amazon forest, corti- caceae, corticolous, 24 May 2011, US (LPB, herb. colous, 25 July 2009, AF 14272 & PR (KRAM, LPB, Schiefelbein 3111). herb. Flakus). *Pallidogramme chrysenteron (Mont.) Staiger, *Graphis subradiata (Nyl.) Lücking Kalb & Lücking Known only from the Neotropics and previ- This species was reported in the Neotropics ously reported from Brazil, Colombia and Costa from Brazil, Colombia, Costa Rica, Guyana and Rica (Lücking et al. 2008, 2009). Panama, and elsewhere from Australia, the Philip- Specimens examined. BOLIVIA. Dept. La Paz. pines and Thailand (Staiger 2002; Wolseley et al. U. SCHIEFELBEIN ET AL.: A CONTRIBUTION TO GRAPHIDACEAE OF BOLIVIA. 2 93

2002; Sipman et al. 2008; Boom & Sipman 2013; (Leight.) Kalb and P. scalpturata; pictures in, e.g., Linsangan-Tabaquero et al. 2013). Lücking et al. 2011; Aptroot & Sparrius 2014), we refer our sample to P. tortuosa. Specimens examined. BOLIVIA. Dept. La Paz. Prov. Bautista Saavedra, Calzada, 15°09′06″S, 68°30′24″W, Specimens examined. BOLIVIA. Dept. La Paz. 835 m, Preandean Amazon secondary forest, corticolous, near Pacallo village, 16°12′10″S, 67°50′39″W, 1360 m, 17 May 2011, MK 8806 (LPB, UGDA). montane forest, corticolous, 3 Aug. 2008, MK 7115 (LPB, UGDA). Phaeographis leprieurii (Mont.) Staiger * pachyspora (Redinger) Staiger Previously known from only a single locality in Bolivia (Kukwa et al. 2013). So far known only from Brazil (Staiger 2002). Specimens examined. BOLIVIA. Dept. La Paz. Specimens examined. BOLIVIA. Dept. La Paz. Prov. Abel Iturralde, Tumupasa village, 14°08′51″S, Prov. Larecaja, near Achiquiri, 15°13′31″S, 68°13′49″W, 67°53′34″W, 350 m, garden in the village, on mango 1071 m, Yungas secondary forest, 16 May 2011, US tree, 31 Aug. 2008, MK 6998 (LPB, UGDA). (LPB, herb. Schiefelbein 2988).

Phaeographis scalpturata (Ach.) Staiger * serpentinellum (Nyl.) Staiger This species was reported from Bolivia before To our knowledge this species was reported by Nylander (1859) and Flakus et al. (2013). only from Colombia (Staiger 2002; Sipman et al. 2008). Specimens examined. BOLIVIA. Dept. Beni. Prov. Ballivian, Palmar village, 14°58′56″S, 67°05′16″W, Specimens examined. BOLIVIA. Dept. La Paz. 250 m, by road, corticolous, 2 Aug. 2008, MK 7109, Prov. Nor Yungas, COTAPATA, Chairo, 16°12′07″S, 7112 (LPB, UGDA); Dept. La Paz. Prov. Nor Yungas, 67°50′05″W, Yungas lower montane forest, 1300 m, Coroico, 16°11′10″S, 67°43′16″W, 1550 m, Yungas corticolous, 21 May 2011, US (LPB). montane forest, corticolous, 6 June 2010, AF 16430.2 & PR (KRAM, LPB); near Pacallo village, 16°12′10″S, * santensis (Tuck.) Kalb 67°50′39″W, 1360 m, montane forest, corticolous, 3 Aug. 2008, MK 7170 (LPB, UGDA); Dept. Santa Cruz. A Pantropical species known in the Neotropics Prov. Guarayos, RN de Vida Silvestre Ríos Blanco y from Brazil, Costa Rica and Paraguay, and else- Negro, Plan de Manejo AISU, 15°01′49″S, 62°46′36″W, where from Australia, Mozambique and the USA 236 m, lowland Amazon forest, corticolous, 25 July (South Carolina) (Frisch 2006; Sipman et al. 2012; 2009, AF 14238, 14242 & PR (KRAM, LPB); ibidem, Esslinger 2014). 15°01′58″S, 62°46′36″W, 242 m, 24 July 2009, AF 13886 & PR (LPB); Virgen de Pilar near Chonta, Specimens examined. BOLIVIA. Dept. Beni. Prov. 15°38′54″S, 62°57′37″W, 229 m, lowland Amazon Cercado, Casa del Tigre near Chuchini, 14°43′17″S, forest, corticolous, 22 July 2009, AF 13257 & PR (LPB). 64°56′53″W, 160 m, lowland Amazon forest, corti- colous, 31 Aug. 2012, MK 11991 (LPB, UGDA). *Phaeographis tortuosa (Ach.) Müll. Arg. Acknowledgements. We are greatly indebted to Rosa Reported so far from the Caribbean, Brazil I. Meneses Q., Director of Herbario Nacional de Bo- and Panama (Staiger 2002; Cáceres 2007; Boom livia, Universidad Mayor de San Andrés, La Paz for & Sipman 2013). her kind hospitality and assistance, to a number of Our material agrees anatomically and chemi- people who were so helpful during the field work, in particular Pamela Rodriguez Flakus (Frankfurt), and cally with the description given by Staiger (2002) to the anonymous reviewers for helpful suggestions on but the thallus is rather smooth (as also pictured the manuscript. This research received support from the by Cáceres 2007) than granular. Taking under con- Polish National Centre for Research and Development sideration the variability of thallus morphology (NCBiR) under the LIDER Programme for 2010–2013 in other Phaeographis species [e.g., P. intricans (No. 92/L-1/09). 94 POLISH BOTANICAL JOURNAL 59(1). 2014

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Received 10 May 2014